MOLLUSCAN ASSEMBLAGES OF RECENT CALCAREOUS TUFAS IN THE PODHALE BASIN AND PIENINY MTS (S. POLAND)

MOLLUSCAN ASSEMBLAGES OF RECENTCALCAREOUS TUFAS IN THE PODHALE BASINAND PIENINY MTS (S. POLAND)

WITOLD PAWE£ ALEXANDROWICZ

Chair of Environmental Analysis, Cartography and Industrial Geology, AGH University of Scienceand Technology, Al. Mickiewicza 30, 30-059 Cracow, Poland (e-mail: [email protected])

ABSTRACT: Recently precipitated travertines and calcareous tufas occur commonly in the Podhale Basin andPieniny Mts. They contain more or less rich subfossil mollusc assemblages which were analysed in 73 samplescollected from 43 localities. Seven types and two subtypes of assemblages were distinguished, according totheir structure and species composition, depending closely on the local ecological conditions. The whole ma-terial comprised 74 species. Only one of them – Bithynella austriaca (Frauenfeld) – was found in many profilesand in substantial numbers, while the remaining species were rare. Malacological investigations of Late Holo-cene and recent calcareous tufas enable a reconstruction of conditions in which such deposits are formed,and determining the diversity of mollusc assemblages associated with the differentiation of the sedimentaryenvironment. They are also useful for actuopalaeontological analysis which is important for palaeogeographicand palaeoecological interpretations of Holocene and Late Glacial calcareous deposits of southern Poland.

KEY WORDS: calcareous tufa, travertine, Late Holocene, molluscan assemblages, Podhale Basin, Pieniny Mts

INTRODUCTION

Calcareous tufa belongs to the most characteristictypes of interglacial sediments. In Poland it was de-posited mainly during the Late Glacial and Holocene.In several geographical regions, especially in the Car-pathians and Middle Polish Uplands, accumulation offresh-water calcareous sediments can be observed alsorecently. These deposits result from precipitation ofcalcium carbonate from water, stimulated by phy-sico-chemical or biological processes, but most oftenas a combination of both. Most localities withEarly/Middle Holocene calcareous tufa and with LateGlacial tufa have been studied in detail usinglithological, palaeontological and isotopic methods.Results of these investigations have been published bynumerous authors (for references see S. W. ALEXAN-DROWICZ 1983, W. P. ALEXANDROWICZ 2004). On theother hand, the youngest sediments, deposited dur-ing the historical period (mainly the last 500 years)and/or recently formed, have not been investigatedin detail till now. Most localities of these sedimentswere found mainly during the geological mapping

and investigations of tectonic structures (MASTELLA1975, GRUSZCZYÑSKI & MASTELLA 1986).

Calcareous tufas show a high diversity controlledby different depositional processes. Two main litho-logical types can be distinguished. There are hard,strongly porous fresh-water limestones (travertines)and granular calcareous tufa composed of blocks oftravertines, calcareous sand, mud and clay. In severallocalities calcareous sediments are accompanied byalluvial sands and muds, slope deposits and peat. Fivetypes of occurrence of such sediments can be distin-guished. One includes calcareous encrustations onpebbles and on base-rock, calcareous clusters, calca-reous encrustations on plants and calcareous covers.The second comprises sediments deposited in small,sometimes temporary water bodies with rich vegeta-tion. Calcareous tufa accumulated on terraces or flatslopes represents the third type. The fourth includescalcareous fans deposited on steep slopes nearsprings or at the base of rock faces. Travertine andgrained calcareous deposits accumulated near water-

FOLIAMALACOLOGICAISSN 1506-7629

The Association of Polish Malacologists

& Faculty of Biology, Adam Mickiewicz University

Poznañ 2010

Vol. 18(3): 99–112

doi:10.2478/v10125-010-0009-9

Unauthenticated | 77.254.129.50Download Date | 7/4/13 10:28 AM

falls constitute the last type (GRUSZCZYÑSKI & MAS-TELLA 1986).

In the Pieniny Mts. and the Podhale Basin hard,strongly porous travertines are most common but in

numerous localities granular tufas are also found.The latter contain more or less rich and diverse mol-lusc assemblages.

MATERIAL AND METHODS

Most calcareous tufas of the Podhale Basin andPieniny Mts are developed as hard travertines andthin (1–5 cm) encrustations or calcareous covers.However, during the field work, granular mollusc--bearing sediments were found in 43 localities (Fig. 1,Table 1). The whole analysed material included 74samples; they were washed to select all identifiablemollusc shells and their fragments. Species identifica-tion was based on monographs, keys and comparativecollections. The frequency was expressed as the num-ber of complete shells and shell fragments, recalcu-lated according to the scheme proposed by LOZEK(1964) and S. W. ALEXANDROWICZ (1987, 1999).More than 8,200 specimens of 74 snail and bivalvespecies were identified (68 land snails, 4 water snails,2 bivalves).

Standard methods of malacological analysis (LO-ZEK 1964, S. W. ALEXANDROWICZ 1987, 1999) were ap-

plied. Each taxon was assigned to one of five ecologi-cal groups: F – shade-loving, forest and shrubland spe-cies, O – open-country species of habitats with variedhumidity (meadow, bare rocks, xerothermic), M –mesophile, species of high ecological tolerance (me-sophile, catholic), H – hygrophile snails and W –aquatic species (of ephemeral and permanent fresh-waters). The results of ecological analysis were pre-sented as triangular diagrams which made it possibleto distinguish seven types and two subtypes of mollus-can assemblages. The taxonomic diversity index(TDI; S. W. ALEXANDROWICZ 1987) was calculated foreach sample. Indices of constancy and dominance(C-D) as well as normalised index of constancy (Ci)and dominance (Di) were also calculated (S. W.ALEXANDROWICZ 1987, 1999). Zoogeographical clas-sification of the fauna followed LOZEK (1964), KÖR-NIG (1966) and S. W. ALEXANDROWICZ (1987, 1999).

LOG DESCRIPTION

During field work fourteen localities of recentlyprecipitating mollusc-bearing calcareous tufa werefound in the Podhale Basin (MP-1–MP-14) and twentynine localities – in the Pieniny Mts (MP-15–MP-43)(Fig. 1, Table 1). Some localities described below hadbeen tentatively mentioned by HALICKI & LILPOP(1932), URBAÑSKI (1932), MASTELLA (1975), S. W.ALEXANDROWICZ (1984a, b), W. P. ALEXANDROWICZ(1997, 2001, 2003, 2004) and others.

Dzianisz (MP-1) – calcareous cover formed ofhard, white travertines, found in the terrace bank ofthe Kijów Stream, the left bank tributary of the Dzia-nisz Stream in Dzianisz village (Fig. 1). Calcareoustufa with numerous travertine blocks was found in asmall niche. A single sample (Mp-1) contained a poormollusc assemblage (8 species and 78 specimens).The fauna was dominated by water taxa: Bithynellaaustriaca (Frauenfeld) and Pisidium personatum Malm(Table 1).

Ostrysz (MP-2) – recently accumulated calcareoustufa forming a small fan near a spring in the middlepart of northern slopes of the Ostrysz Hill (Fig. 1).This fan was twenty centimetres thick and built ofcoarse- and fine-grained calcareous sediment withsmall fragments of strongly porous, white travertines.Two samples (Mp-2 and Mp-3) contained a relatively

rich fauna, composed of 22 species and 893 speci-mens (Table 1). Bithynella austriaca (Frauenfeld)clearly prevailed, constituting more than 90% of speci-mens. The large exposure of Late Glacial, Early andMiddle Holocene calcareous tufa, situated in the up-per part of this slope, has been described in detail byW. P. ALEXANDROWICZ (1997, 2001).

Gliczarów (MP-3) – young calcareous sedimentsaccumulated near a small waterfall about 50 m belowthe road from Czarny Dunajec to Lower Gliczarów(Fig. 1). They were developed as yellowish-gray granu-lar tufa, about 30 cm thick, consisting of numeroustravertine blocks and underlain by a single layer ofwhite, hard travertine. Four samples (Mp-3–Mp-7)were collected in this locality (Table 1). Each con-tained a poor fauna with many shells of Bithynellaaustriaca (Frauenfeld). This dominant species was ac-companied by snails of moist, more or less shady habi-tats (Carychium minimum O. F. Müller, Zonitoides niti-dus (O. F. Müller), Perforatella vicina (Rossmässler)),as well as by catholic species (Cochlicopa lubrica (O. F.Müller) and Euconulus fulvus (O. F. Müller)). In all, 19taxa and 371 shells were found in the locality (Table1). The main exposure of Late Glacial, Early andMiddle Holocene calcareous tufa in Gliczarów hasbeen described by several authors (HALICKI & LILPOP

100 Witold Pawe³ Alexandrowicz


1932, URBAÑSKI 1932, PAZDUR 1987, W. P.ALEXANDROWICZ 1997, 2003 and others) is situatedabout 100 m higher, on the slope of the GliczarowskiWierch.

Czarna Góra (MP-4) – coarse-grained calcareoustufa with blocks of hard travertines, forming an irreg-ular lens within alluvial gravel and sand. The de-scribed locality is situated on the flat part of the land-slide developed on the western slope of Czarna Góranear Bia³ka Tatrzañska (Fig. 1). A single sample(Mp-8) contained more than 200 shells representing18 species (Table 1). The fauna was dominated by twogroups of molluscs: shade-loving taxa preferringmore or less moist habitats: Vestia gulo (E. A. Bielz),Perforatella vicina (Rossmässler), Vitrea diaphana(Studer), Vitrea crystallina (O. F. Müller), and numer-ous specimens of Bithynella austriaca (Frauenfeld), ac-companied by single shells of Galba truncatula (O. F.Müller).

Trybsz (MP-5) – calcareous encrustations precipi-tated on the moss near a spring situated on the lowerterrace of the Trybska River in Trybsz village (Fig. 1).A poor mollusc assemblage (3 species and 111 speci-mens), composed exclusively of water species, mainlyBithynella austriaca (Frauenfeld) and Pisidiumpersonatum Malm, was found there (sample Mp-9)(Table 1).

£apsze Wy¿ne (MP-6) – calcareous cover with sev-eral niches filled by granular calcareous tufa, locatednear a small spring on the eastern slope of the BliŸnaHill. Its mollusc assemblage was relatively rich andcontained 16 species represented by 125 specimens(sample Mp-10). Bithynella austriaca (Frauenfeld) wasthe most important component of this fauna. Hygro-phile and mesophile taxa completed the assemblage,while shade-loving ones were few (Table 1). The local-ity was first mentioned by W. P. ALEXANDROWICZ(1997).

£apszanka (MP-7) – granular calcareous tufa andtwo layers of hard, porous travertines located withinsediments forming the lower terrace of the £apszankaStream (Fig. 1). The fauna included 15 species andnearly 200 specimens (sample Mp-11). Watermolluscs (Bithynella austriaca (Frauenfeld) andPisidium personatum Malm), snails inhabiting forests(Perforatella vicina (Rossmässler), Aegopinella pura (Al-der)) and catholic taxa (Cochlicopa lubrica (O. F.Müller)) were the main components of this assem-blage (Table 1).

Stawiska (MP-8) – fine- and coarse-grained calcare-ous tufa abounding in blocks of hard travertines, de-veloped as a fan on the flat surface of the lower ter-race of the Kacwiñski Stream valley, near the southernborder of the Stawiska hamlet (Fig. 1). The thicknessof this fan is up to 50 cm, and the calcareous sedi-ments are underlain by alluvial gravel. In two samples(Mp-12 and Mp-13) a very rich and diverse mollusc as-semblage (total of 38 species and nearly 1,000 speci-

mens) was found (Table 1). Water molluscs: Bithynellaaustriaca (Frauenfeld), Pisidium personatum Malm, Gal-ba truncatula (O. F. Müller) and mesophile snails:Cochlicopa lubrica (O. F. Müller), Euconulus fulvus (O.F. Müller), Punctum pygmaeum (Draparnaud), andsome other taxa were the most important ecologicalgroups of this fauna. Forest and shade-loving snails:Acicula polita (Hartmann), Eucobresia nivalis (Dumontet Mortillet) and hygrophile species – Carychium mini-mum (O. F. Müller) supplemented this assemblage.Single shells of open-country meadow species: Pupillamuscorum (Linnaeus) and Vallonia pulchella (O. F. Mül-ler), were also noted. This locality was first describedby W. P. ALEXANDROWICZ (1997).

£apsze Ni¿ne (MP-9) – calcareous encrustationson moss, precipitating near a spring situated on a lowterrace of a small stream close to the outlet to the£apszanka river on the eastern border of the village£apsze Ni¿ne (Fig. 1). Quite a poor fauna (4 species,43 shells), clearly dominated by Bithynella austriaca(Frauenfeld), was found in the single sample (Mp-14)collected in this locality (Table 1).

Kacwin I (MP-10) – calcareous cover with nichesfilled by granular calcareous tufa was found near thelower waterfall in the Kacwinka Stream valley (Fig. 1).In the single sample (Mp-15) only four species and 68specimens were found, forming an assemblage withmore then 90% of Bithynella austriaca (Frauenfeld)(Table 1).

Kacwin II (MP-11) – calcareous encrustations onplants near the upper waterfall in the KacwinkaStream valley (Fig. 1). A poor molluscan assemblage(4 species and 28 specimens) found in this place wasquite similar to the one described above (sampleMp-16) (Table 1).

Niedzica (MP-12) – coarse-grained calcareous tufawith large fragments of hard travertines, deposited ina small stream valley in the eastern part of the Niedzi-ca village (Fig. 1). Two samples (Mp-17 and Mp-18)contained a relatively rich fauna (23 taxa and 141specimens) (Table 1). Shade-loving species: Vitrea su-brimata (Reinhardt), Perforatella incarnata (O. F.Müller), Isognomostoma isognomostoma (Schröter) andfew others were the most important components ofthis assemblage. Mesophile and hygrophile species aswell as water molluscs completed the fauna. A profileof Middle Holocene calcareous tufa containing a veryrich fauna with forest snails, located about 100 m westof this site, was described by W. P. ALEXANDROWICZ(1997).

Niedzica Majerz (MP-13) – fine-grained calcareoustufa, accumulated in the middle part of southernslopes of the Majerz Hill, close to the spring in a smalland much vegetated kettle (Fig. 1). A poor fauna (6taxa and 22 specimens) dominated by hygrophile spe-cies: Zonitoides nitidus (O. F. Müller) and Succineaputris (Linnaeus) was found in a single sample(Mp-19) (Table 1).

Molluscan assemblages of tufas in the Podhale Basin and Pieniny Mts 101


Falsztyn (MP-14) – granular calcareous tufa withfew fragments of hard travertines and numerousblocks of limestones, forming a fan at the foot of rockface of the Patria Rocky Hill in Falsztyn (Fig. 1). Twocollected samples (Mp-20 and Mp-21) contained avery rich and diverse mollusc fauna (35 taxa and morethan 350 specimens) (Table 1). Shade-loving species:Vitrea crystallina (O. F. Müller), V. diaphana (Studer),Aegopinel la pura (Alder), Arianta arbustorum(Linnaeus), Perforatella incarnata (O. F. Müller) andmany others were the main components of this assem-blage. Catholic species: Nesovitrea hammonis (Ström),Cochlicopa lubrica (O. F. Müller) and Carychium triden-tatum (Risso), constituted the second most importantgroup, while hygrophile and water species occurredrarely.

Czorsztyn (MP-15) – granular calcareous tufa 10cm thick cropping out close to the spring near theCzorsztyn castle (Fig. 1). A poor molluscan fauna (11taxa and 65 specimens) with numerous shells of Bithy-nella austriaca (Frauenfeld) accompanied by catholicsnails was found in sample Mp-22 (Table 1).

Harcygrunt (MP-16) – calcareous tufa found by S.W. ALEXANDROWICZ (1984a) in the HarcygruntStream valley near a spring at the foot of the largelandslide (Fig. 1). In one sample (Mp-23) shells repre-senting all ecological groups of molluscs (13 speciesbut only 25 specimens) were found (Table 1).

Limbargowy Stream (MP-17) – coarse-grained cal-careous tufa forming a layer about 20 cm thick, distin-guished in top part of the lower terrace in the Limbar-gowy Stream valley. Its mollusc assemblage (13 spe-cies, 123 specimens) found in two samples (Mp-24,Mp-25) was dominated by forest snails: Trichia uniden-tata (Draparnaud), Perforatella vicina (Rossmässler),Arianta arbustorum (Linnaeus) as well as by catholicspecies (Cochlicopa lubrica (O. F. Müller), Euconulusfulvus (O. F. Müller) and Nesovitrea hammonis(Ström)) (Table 1).

K¹ty (MP-18) – fine-grained calcareous tufa in asmall marsh, strongly overgrown by plants. Thismarsh is situated near a spring, on the flat surface ofthe lower terrace (Fig. 1). In two samples (Mp-26,Mp-27) collected here, a rich and diverse molluscan


Fig. 1. Localisation of profiles of Upper Holocene calcareous tufa in Podhale Basin and Pieniny Mts. 1 – towns and villages,2 – Czorsztyn dam lake, 3 – main sites, 4 – supplementary sites, 5 – state border, 6 – rivers and streams



Table 1. Profiles of Upper Holocene calcareous tufa in Podhale Basin and Pieniny Mts. Ntax – number of taxa, Nspec – num-ber of specimens

Locality Samples Ntax Nspec

MP-1 Dzianisz Mp-1 8 78

MP-2 Ostrysz Mp-2, Mp-3 22 893

MP-3 Gliczarów Mp-4 – Mp-7 19 371

MP-4 Czarna Góra Hill Mp-8 18 206

MP-5 Trybsz Mp-9 3 111

MP-6 £apsze Wy¿ne Mp-10 16 125

MP-7 £apszanka Mp-11 15 196

MP-8 Stawiska Mp-12, Mp-13 38 997

MP-9 £apsze Ni¿ne Mp-14 4 43

MP-10 Kacwin I Mp-15 4 68

MP-11 Kacwin II Mp-16 4 28

MP-12 Niedzica Mp-17, Mp-18 23 141

MP-13 Majerz Mp-19 6 22

MP-14 Falsztyn Mp-20, Mp-21 35 352

MP-15 Czorsztyn Mp-22 11 64

MP-16 Harcygrunt Mp-23 13 25

MP-17 Limbargowy Stream Mp-24, Mp-25 13 123

MP-18 K¹ty Mp-26, Mp-27 34 424

MP-19 Straszny Stream Mp-28 – Mp-30 12 42

MP-20 Macelowy Stream Mp-31 20 103

MP-21 Kot³owy Gorge Mp-32, Mp-33 13 151

MP-22 Gorczyñski Gorge Mp-34 18 159

MP-23 Sobczañski Gorge Mp-35 9 71

MP-24 Pieniñski Stream I Mp-36, Mp-37 23 241

MP-25 Pieniñski Stream II Mp-38, Mp-39 8 167

MP-26 Kroœcienko Toporzyska I Mp-40, Mp-41 17 70

MP-27 Kroœcienko Toporzyska II Mp-42, Mp-43 17 65

MP-28 Ociemny Stream Mp-44 22 273

MP-29 D³ugi Gronik Mp-45 – Mp-48 22 288

MP-30 Szczawny Stream Mp-49, Mp-50 11 227

MP-31 Zakijowski Stream Mp-51, Mp-52 23 226

MP-32 Skotnicki Stream I Mp-53, Mp-54 19 86

MP-33 Skotnicki Stream II Mp-55, Mp-56 19 43

MP-34 Skotnicki Stream III Mp-57 – Mp-59 25 316

MP-35 G³êboki Stream Mp-60 10 83

MP-36 Grajcarek Stream I Mp-61 – Mp-63 29 181

MP-37 Grajcarek Stream II Mp-64, Mp-65 14 95

MP-38 Pa³kowski Stream Mp-66 4 83

MP-39 Krupianka Mp-67, Mp-68 27 133

MP-40 Szlachtowa Mp-69, Mp-70 10 130

MP-41 Homole Gorge Mp-71, Mp-72 29 308

MP-42 Zaskale Mp-73 5 174

MP-43 Bia³a Woda Gorge Mp-74 14 364


assemblage with 34 species and 424 specimens wasfound. Hygrophile taxa, mainly Succinea putris (Lin-naeus), were the most important components of thefauna. Water molluscs, represented by Bithynellaaustriaca (Frauenfeld) and Galba truncatula (O. F.Müller), catholic taxa like Nesovitrea hammonis(Ström), Cochlicopa lubrica (O. F. Müller) and forestsnails: Vitrea subrimata (Reinhardt), Isognomostoma iso-gnomostoma (Schröter) and Chilostoma faustinum(Rossmässler), were also numerous. Single shells ofmeadow species were present (Table 1).

Straszny Stream (MP-19) – travertine cover withsmall niches filled by granular calcareous tufa con-taining fragments of travertines, cropping out nearthe spring in the middle part of the Straszny Streamvalley (Fig. 1). A poor mollusc fauna (12 species and42 specimens) was found in three samples (Mp-28,Mp-29 and Mp-30). Mesophile snails representedmainly by Succinea oblonga Draparnaud were the mostimportant components of the assemblage; they wereaccompanied by water molluscs and shade-loving spe-cies (Table 1).

Macelowy Stream (MP-20) – thin lens of granularcalcareous tufa cropping out close to the small springin the lower course of the Macelowy Stream valley (Fig.1). Relatively rich fauna (20 taxa and more than 100specimens) was found in a single sample (Mp-31). Wa-ter species: Bithynella austriaca (Frauenfeld.) and Galbatruncatula (O. F. Müller) as well as snails living in moisthabitats: Carychium minimum O. F. Müller, Vertigoantivertigo (Draparnaud) were the main componentsof the assemblage, while specimens of the remainingecological groups were few (Table 1). Several metresdownstream from this locality, a Late Holocene calcar-eous mollusc-bearing tufa, radicarbon-dated at1,430±60 (Gd-5112) and 900±50 (Gd-5259) years BPwas described by S. W. ALEXANDROWICZ (1990).

Kot³owy Gorge (MP-21) – a small outcrop ofcoarse-grained calcareous tufa with numerous blocksof hard travertines and angular fragments of lime-stones was found near a spring in the upper part ofthe Kot³owy Gorge (Fig. 1). In two samples (Mp-32and Mp-33) a poor fauna of molluscs (13 taxa, 151specimens) was found (Table 1). The high proportionof water snails: Bithynella austriaca (Frauenfeld) andGalba truncatula (O. F. Müller), was the most charac-teristic feature of this assemblage, while species of theremaining ecological groups were represented occa-sionally.

Gorczyñski Gorge (MP-22) – calcareous cover withnumerous blocks of limestones cropping out in theuppermost part of the Gorczyñski Gorge (Fig. 1).Sample Mp-34 contained a rich fauna (18 species and159 specimens) dominated by water molluscs, mainlyBithynella austriaca (Frauenfeld) and Pisidium perso-natum Malm. Snails typical of shaded habitats, likeVitrea diaphana (Studer) and Perforatella vicina (Ross-mässler) were additional components of the fauna,

while the remaining ecological groups were subordi-nate (Table 1). This locality was first noted by S. W.ALEXANDROWICZ (1984b).

Sobczañski Gorge (MP-23) – small fan of recentlyaccumulated sediments situated at the foot of therock face in the Sobczañski Gorge close to its gate.The fan was composed of fine-grained calcareous tufaabounding in limestone fragments (Fig. 1). A poorfauna (9 species and 71 specimens) with numerousshells of Bithynella austriaca (Fauenrfeld) was found insample Mp-35 (Table 1). Other localities of young cal-careous tufa in the Sobczañski Gorge were describedby S. W. ALEXANDROWICZ (1984b, 1988), S. W.ALEXANDROWICZ et al. (1985) and W. P. ALEXANDRO-WICZ (2004).

Two localities of granular calcareous tufa areknown in the Pieniñski Stream valley. One of them –Pieniñski Stream I (MP-24), a layer of granular calca-reous tufa with irregular fragments of travertines andlimestones 15 cm thick, was found near a spring in themiddle part of the Pieniñski Stream valley (Fig. 1).Two samples taken one below another (Mp-36,Mp-37) yielded a relatively rich fauna – 23 species andalmost 250 specimens (Table 1). In the lower sample(Mp-35) a high proportion of shade-loving species:Isognomostoma isognomostoma (Schröter), Perforatella vi-cina (Rossmässler), Arianta arbustorum (Linnaeus),Aegopinella pura (Alder) and others was noted. The as-semblage was supplemented by the following catholicsnails: Carychium tridentatum (Risso), Cochlicopa lubrica(O. F. Müller) and Euconulus fulvus (O. F. Müller).Species of the remaining ecological groups occurredrarely. The mollusc assemblage in the second samplewas poorer and dominated by water and mesophilesnails: Bithynella austriaca (Frauenfeld), Cochlicopa lub-rica (O. F. Müller) and Nesovitrea hammonis (Ström).

Pieniñski Stream II (MP-25) – the locality was situ-ated 100–150 m upstream of the preceding one. Gra-nular calcareous tufa occurred there in two niches de-veloped within a travertine cover (Fig. 1). Twosamples (Mp-38, Mp-39) contained a poor fauna (8species, 167 specimens) dominated by water molluscs:Bithynella austriaca (Frauenfeld) and Pisidium perso-natum Malm (Table 1).

Two sites of hard travertines with niches filling bygranular calcareous tufa were found in a small streamvalley south-west of the centre of Kroœcienko (Kroœ-cienko–Toporzyska). In the first one – Kroœcienko-To-porzyska I (MP-26) – two layers of white, hard andstrongly porous travertines separated by granular cal-careous tufa occurred (Fig. 1). Two samples (Mp-40,Mp-41) contained a relatively poor fauna (17 species70 specimens), including mainly snails of shaded habi-tats and mesophile snails: Oxychilus depressus (Sterki),Laciniaria plicata (Draparnaud) and Euconulus fulvus(O. F. Müller).

Kroœcienko-Toporzyska II (MP-27) – very similar tothe preceding one, situated about 70 m upstream



(Fig. 1). Calcareous sediment in two samples (Mp-42,Mp-43) contained a fauna of 17 taxa represented by65 shells, with a high proportion of water molluscs(Table 1).

Ociemny Stream (MP-28) – calcareous tufa in theform of a small fan near a spring in the middle part ofthe Ociemny Stream valley (Fig. 1). A rich fauna wasfound in sample Mp-44, with 22 species and almost300 specimens. The community comprised two maingroups of molluscs: one with Bithynella austriaca(Frauenfeld) and Galba truncatula (O. F. Müller), an-other represented by snails of more or less shadedmoist habitats: Succinea putris (Linnaeus), Vestia gulo(E. A. Bielz.) and Perforatella vicina (Rossmässler).Catholic and forest species supplemented this assem-blage. Several metres downstream a locality of Lowerand Middle Holocene mollusc-bearing deposits wasfound and described in detail by W. P. ALEXANDRO-WICZ (2004).

D³ugi Gronik (MP-29) – calcareous sedimentsabout 50 cm thick, developed near a spring as hardporous travertines, forming two layers separated bycoarse-grained tufa (Fig. 1). Four samples (Mp-45– Mp-48) were collected in this site. They contained arelatively poor mollusc assemblage (22 taxa andnearly 300 specimens). The fauna was dominated bycatholic snails: Nesovitrea hammonis (Ström), Cochli-copa lubrica (O. F. Müller), Succinea oblonga Drapar-naud and others. Molluscs of the remaining ecologi-cal groups were few (Table 1).

Szczawny Stream (MP-30) – travertine fan withsmall niches filled with granular calcareous tufa, lo-cated in the middle part of the Szczawny Stream valleynorth-east of Kroœcienko (Fig. 1). Two samples(Mp-49 and Mp-50) contained 227 specimens of 11species. Two groups of molluscs were the most impor-tant components of this fauna. Water species, mainlyBithynella austriaca (Frauenfeld) and Pisidium persona-tum Malm, constituted the first one, while snails typi-cal of moist, more or less shaded habitats: Perforatellavicina (Rossmässler) and Zonitoides nitidus (O. F.Müller), belonged to the second group (Table 1).

Zaki jowski Stream (MP-31) – outcrop ofcoarse-grained calcareous tufa, located in the middlepart of the Zakijowski Stream valley, on the lower ter-race about 10 m below a small spring (Fig. 1). Thewhole collected material (samples Mp-51 and Mp-52)included 23 species and 226 specimens. Hygrophilesnails: Carychium minimum O. F. Müller and Succineaputris (Linnaeus), as well as species inhabiting moistforests: Macrogastra ventricosa (Draparnaud), Vestiagulo (E. A. Bielz.) and Perforetella vicina (Rossmässler),were the most important components of this fauna. Inthe upper sample (Mp-52) numerous shells of watermolluscs: Galba truncatula (O. F. Müller) and Pisidiumpersonatum Malm were also noted. Snails typical ofshaded habitats and catholic species supplementedthe community (Table 1).

Three localities of recently deposited calcareoustufa were found in the Skotnicki Stream valley (north-ern part of Szczawnica). In the outcrop SkotnickiStream I (MP-32), situated about 300 m upstream ofthe stream outlet (Fig. 1), calcareous tufa was devel-oped as a thin layer between alluvial deposits. Sam-ples Mp-53 and Mp-54 contained a relatively poorfauna (19 taxa, 86 specimens) with a high proportionof Bithynella austriaca (Frauenfeld), accompanied byforest and catholic species (Table 1).

Skotnicki Stream II (MP-33) – the second localityis about 100 m upstream of the preceding one. Cal-careous sediments contained a poor fauna (19 spe-cies, 43 specimens) found in samples Mp-55 andMp-56. The assemblage was dominated by forestsnails: Arianta arbustorum (Linnaeus) and Aegopinellapura (Alder), with a limited proportion of species rep-resenting the remaining ecological groups (Table 1).

Skotnicki Stream III (MP-34) – granular calcareoustufa up to 20 cm thick cropping out on the terrace inthe upper part of the Skotnicki Stream valley (Fig. 1).Three samples: Mp-56, Mp-57 and Mp-58, containinga fauna with 25 taxa and 315 specimens, were col-lected in this locality. In the lower part of the layer thefauna was dominated by forest snails: Vitrea trans-sylvanica (Clessin), Arianta arbustorum (Linnaeus),Discus perspectivus (Mühlfeld). The assemblage foundin the uppermost sample (Mp-58) included numer-ous shells of Bithynella austriaca (Frauenfeld), whichconstituted about 90% of the assemblage (Table 1).

G³êboki Stream (MP-35) – calcareous encrustationson plants, found near a small spring at the mouth partof the G³êboki Stream valley in the southern part ofSzczawnica (Fig. 1). Only one sample (Mp-60), com-prising a very poor mollusc assemblage, derives fromthis locality. The most important component was Bithy-nella austriaca (Frauenfeld), accompanied by forestsnails: Vitrea diaphana (Studer), Oxychilus glaber (Ross-mässler) and Vestia gulo (E. A. Bielz) (Table 1).

Two localities of recently deposited, coarse-grain-ed calcareous tufa were located in the GrajcarekStream valley between Szczawnica and Szlachtowa. Inthe site Grajcarek Stream I (MP-36) a layer of granu-lar calcareous tufa 15 cm thick cropped out in the up-per part of a low terrace of the Grajcarek Stream (Fig.1). Three samples (Mp-61, Mp-62 and Mp-63) con-tained quite a rich mollusc assemblage with 29 speciesand 181 specimens. Forest snails: Aegopinella pura (Al-der), Eucobresia nivalis (Dumont et Mortillet) andOxychilus depressus (Sterki) were frequent, while catho-lic species and water molluscs: Vitrea contracta(Westerlund) and Bithynella austriaca (Frauenfeld)supplemented the fauna (Table 1).

Grajcarek Stream II (MP-37) – a locality very simi-lar to the preceding one, situated near the outlet ofthe Sopotnicki Stream (Fig. 1). Samples Mp-64,Mp-65 contained a poor fauna (14 species and lessthan 100 specimens) with Bithynella austriaca (Frauen-



feld), forest snails – Aegopinella pura (Alder) andArianta arbustorum (Linnaeus) as well as hygrophilespecies – Succinea putris (Linnaeus). A few other out-crops of alluvial sediments with mollusc-bearing lay-ers of granular calcareous tufa, located in the Graj-carek Stream valley between Szlachtowa and Jaworki,were described by W. P. ALEXANDROWICZ (2004).

Pa³kowski Stream (MP-38) – travertine cover with asmall niche filled with coarse-grained calcareous tufa,deposited near a spring in the middle part of thePa³kowski Stream valley (Fig. 1). A very poor molluscfauna (4 species and 83 specimens) was found insample Mp-66. It was dominated by Bithynella austriaca(Frauenfeld) which constituted nearly 90% of thewhole assemblage (Table 1).

Krupianka (MP-39) – thin layer (20 cm) ofcoarse-grained calcareous tufa with numerous frag-ments of hard travertines, found on the surface of thelower terrace of the Krupianka Stream (Fig. 1). It con-tained a rich fauna (27 species, 133 specimens) ex-tracted from two samples (Mp-67, Mp-68). A high pro-portion of forest snails: Aegopinella pura (Alder) andEucobresia nivalis (Dumont et Mortillet), accompaniedby catholic species: Euconulus fulvus (O. F. Müller)and Cochlicopa lubrica (O. F. Müller), was the mostcharacteristic feature of the assemblage (Table 1).

Szlachtowa (MP-40) – travertine cover with nichesfilled by coarse-grained calcareous tufa containingfragments of travertines, cropping out close to aspring near the outlet of the Sielski Stream valley inthe Szlachtowa village (Fig. 1). A relatively poor fauna(10 species, 130 specimens) with numerous shells ofBithynella austriaca (Frauenfeld) was found in samplesMp-69 and Mp-70 (Table 1).

Homole Gorge (MP-41) – granular calcareous tufawith numerous blocks of hard travertines and frag-ments of limestones, forming a cone near the en-trance to the Homole Gorge (Fig. 1). A rich and di-verse malacofauna (29 taxa and more than 300 speci-mens) was found in samples Mp-71 and Mp-72.Shade-loving species were the most important compo-nents of this assemblage. There were both typical for-est snails: Vitrea diaphana (Studer), Aegopinella pura(Alder), Macrogastra plicatula (Draparnaud), andsnails associated mainly with moist forests, such as:

Macrogastra ventricosa (Draparnaud), Vestia gulo (E. A.Bielz), V. turgida (Rossmässler), Perforatella vicina(Rossmässler). Shells of catholic species: Euconulusfulvus (O. F. Müller), Cochlicopa lubrica (O. F. Müller)and Punctum pygmaeum (Draparnaud), were also nu-merous. Species of other ecological groups occurredoccasionally (Table 1).

Zaskale (MP-42) – calcareous encrustations onplants near a small spring in the upper course of theSkalski Stream valley (Fig. 1). Sample Mp-73 con-tained a very a poor fauna: 5 species and 174 speci-mens. Two water taxa: Bithynella austriaca (Frauen-feld) and Pisidium personatum Malm constituted about85% of this assemblage (Table 1). Numerous local-ities of Holocene calcareous tufa in the SkalskiStream valley were recorded by S. W. ALEXANDRO-WICZ (1984b, 1985) and described in detail by W. P.ALEXANDROWICZ (2004).

Bia³a Woda Gorge (MP-43) – small fan situated atthe foot of the rock face in the middle part of theBia³a Woda Gorge. The fan is built of fine-grained cal-careous tufa with very numerous of fragments of lime-stones (Fig. 1). The fauna in sample Mp-74 was com-posed of 14 species and 364 specimens. Bithynellaaustriaca (Frauenfeld) was the most typical compo-nent of this assemblage. It was accompanied by opencountry snails of dry, sunny habitats: Pupilla muscorum(Linnaeus), Vallonia pulchella (O. F. Müller) and V.costata (O. F. Müller). Species associated with shadedhabitats occurred rarely, while catholic and hygro-phile snails were practically absent (Table 1).

Supplementary sites included numerous other lo-calities of Late Holocene and recently accumulatedmollusc-bearing calcareous tufa from the PieninyMts., described by S. W. ALEXANDROWICZ (1985,1993, 1996) (Witów, Œcigocki Stream, Koz³eckiStream) and W. P. ALEXANDROWICZ (2004) (Tylka,P³aœnie, Zawiasy, Klimontowski Stream, Koz³eckiStream) (Fig. 1). In some of them the deposits wereradiocarbon-dated: Œcigocki Stream – 250±40 BP(Gd-3095) and 640±50 BP (Gd-1807), Koz³eckiStream – 140±50 BP (Gd-5754) (S. W. ALEXANDRO-WICZ 1985, 1993, 1996), Tylka – 535±30 BP (Gd-3197),P³aœnie – 680±80 BP (Gd-6268) (W. P. ALEXANDRO-WICZ 2004).

RESULTS

The outcrops and profiles of Late Holocene and re-cently accumulated calcareous tufa, described above,comprise very diverse molluscan assemblages. Theirtaxonomic composition and structure are closely asso-ciated with the depositional environments. The num-ber of taxa in particular samples varies from 4 to 35.Poor assemblages include less than 15 species per sam-ple while rich ones – between 15 and 35. A great pro-portion (ca. 72%) of the whole analysed material rep-

resents the first group while rich and diverse commu-nities were found in 20 samples (28%) (Fig. 2). Thenumber of specimens varies from 4 to 708 per sampleand in most assemblages it is generally 40–200 speci-mens (60% of samples). Both poorer (4–40 speci-mens) and richer (more than 200 specimens) assem-blages were found in about 20% of the samples (Fig. 2).

All the ecological groups of molluscs defined byLOZEK (1964) and S. W. ALEXANDROWICZ (1987, 1999)



are represented in the analysed material. The quantita-tive relationships between these groups in particularsamples are used to characterise the diversity of assem-blages. Seven types and two subtypes of assemblagescan be distinguished on this basis (Figs 3–5).

AW – assemblage with water molluscs. A typical fea-ture of this usually poor assemblage is the very com-mon occurrence of Bithynella austriaca (Frauenfeld).It is a stenoecious snail, inhabiting mainly springareas of stable temperature. Several species of the ge-nus have been distinguished as a result of the revisionby FALNIOWSKI (1987). Their identification based onshell alone is practically impossible. Thus, in thisstudy, the taxon is understood in its broad sense andincludes the few taxa distinguished as a result of therevision. On the other hand, all species of the genusBithynella, recorded from Poland, occur in similarhabitats and often together. From the point of view ofpalaeoecological interpretation their identification isnot a major issue. B. austriaca often constitutes from50 to 80% of the total number of specimens in asample and is the dominant component of the assem-blage. It is often accompanied by Pisidium personatumMalm and Galba truncatula (O. F. Müller). The firstmentioned taxon inhabits the vicinity of springs andfast flowing mountain streams. The second species istypical of small and much vegetated ponds and tem-porary water bodies. Other water molluscs, such asAncylus fluviatilis O. F. Müller, Radix peregra (O. F.Müller) and Pisidium casertanum (Poli) occur occa-sionally. The limited admixture of land snails prefer-ring moist habitats: Succinea putris (Linnaeus), Per-foratella vicina (Rossmässler), Vestia gulo (E. A. Bielz)is usually noted. The remaining ecological groupsoccur rarely. The assemblage with a high proportionof Bithynella austriaca (Frauenfeld) is most typical ofcalcareous tufa deposited close to springs (S. W.ALEXANDROWICZ 1984b, W. P. ALEXANDROWICZ2004). The described type of fauna was found in 26samples (36% of the total number) (Figs 3, 5).

AW-FM – assemblage with water molluscs, catholicand shade-loving snails. It comprises three groups ofspecies. Water molluscs, constituting about half of theassemblage, are represented mainly by Bithynellaaustriaca (Frauenfeld) and Galba truncatula (O. F.Müller). Species of moist forests: Perforatella vicina(Rossmässler), Vestia gulo (E. A. Bielz) and addition-ally typical woodland snails represent the secondgroup, while catholic species, such as Cochlicopalubrica (O. F. Müller), Euconulus fulvus (O. F. Müller),Carychium tridentatum (Risso) and few others supple-ment this assemblage. A limited admixture of hygro-phile taxa (Succinea putris (Linnaeus)) was also noted.The described assemblage, found in six samples(about 8%) is typical of forests of moist valley bottomswith numerous springs (Figs. 3, 5).

AFM – assemblage with woodland and mesophilesnails. It is a very rich and diverse assemblage, found

in calcareous tufa accumulated within forests, and re-corded in 27 samples (37% of the total number) (Figs3, 5). Two sub-types of this assemblage can be distin-guished.

The sub-assemblage with forest species (AF) rep-resents a rich fauna including typical woodlandsnails, represented by numerous species, such asVitrea diaphana (Studer), Aegopinella pura (Alder),Macrogastra plicatula (Draparnaud), Vitrea crystallina


Fig. 2. Relation between number of taxa (Ntax) and numberof specimens (Nspec) in Upper Holocene calcareous tufain Podhale Basin and Pieniny Mts

Fig. 3. Mollusc assemblages of Upper Holocene calcareoustufa in Podhale Basin and Pieniny Mts. Ecological groupsof molluscs (according to LOZEK 1964, S. W. ALEXAN-DROWICZ 1987, 1999): W – water species, H – hygrophilespecies, M – mesophile species, F – forest species. Forsymbols of assemblages see text


(O. F. Müller), Arianta arbustorum (Linnaeus),Perforatella incarnata (O. F. Müller) and others. Theyare accompanied by taxa of shaded and moist habi-tats: Perforatella vicina (Rossmässler), Vestia gulo (E. A.Bielz) as well as by mesophile snails: Euconulus fulvus(O. F. Müller), Cochlicopa lubrica (O. F. Müller) andPunctum pygmaeum (Draparnaud). In many samplesnumerous shells of Bithynella austriaca (Frauenfeld)are also present. Hygrophile species: Succinea putris(Linnaeus) and Zonitoides nitidus (O. F. Müller) aresubordinate components of this fauna, whileopen-country taxa are practically absent. This faunacharacterises profiles of calcareous tufa situated onthe surface of lower terraces developed in relativelynarrow and much wooded stream valleys. The sub-as-

semblage in question was found in 17 samples (23%)(Figs 4, 5).

The sub-assemblage with mesophile species (FM)comprises catholic species with a substantial admix-ture of shade-loving ones. The former are usually rep-resented by numerous shells of Euconulus fulvus (O. F.Müller), Cochlicopa lubrica (O. F. Müller), Punctumpygmaeum (Draparnaud), Nesovitrea hammonis (Ström)and other species of wide ecological tolerance. Snailsinhabiting mainly sparse forests and shrubland, suchas Arianta arbustorum (Linnaeus), Vitrea crystallina (O.F. Müller), as well as those living in moist forests:Perforatella vicina (Rossmässler) and Vestia gulo (E. A.Bielz) also occur commonly. The fauna is supple-mented by water and hygrophile molluscs. The de-scribed fauna is characteristic of calcareous tufa de-posited on bush-covered terraces. This sub-assembla-ge was found in 10 samples (14%) (Figs 4, 5).

AFM-H – Assemblage with shade-loving and hygro-phile snails. It is a relatively rich assemblage with ahigh proportion of two ecological groups. One com-prises woodland species typical for moist, shaded habi-tats: Perforatella vicina (Rossmässler), Vestia gulo (E. A.Bielz), Perforatella umbrosa (C. Pfeiffer), Macrogastraventricosa (Draparnaud) as well as catholic species:Euconulus fulvus (O. F. Müller), Cochlicopa lubrica (O.F. Müller) and others. Hygrophile snails: Succineaputris (Linnaeus), Zonitoides nitidus (O. F. Müller),Carychium minimum O. F. Müller, belong to the othergroup. Water species, represented mainly by Galbatruncatula (O. F. Müller) and also Bithynella austriaca(Frauenfeld), occur commonly. This fauna occurs incalcareous tufa accumulated in marshes, on flat sur-faces of lower bush-covered terraces. The fauna wasfound in five samples (7%) (Figs 3, 5).

AFH – assemblage of hygrophile species. It is apoor assemblage dominated by species of moist, moreor less open habitats: Succinea putris (Linnaeus),Zonitoides nitidus (O. F. Müller), Carychium minimumO. F. Müller and Vertigo antivertigo (Draparnaud). Wa-ter molluscs typical of temporary water bodies: Galbatruncatula (O. F. Müller) and snails inhabiting moistforests – Perforatella vicina (Rossmässler) – supplementthis community. Species of the remaining ecologicalgroups occur rarely. This fauna is typical of calcareoustufa developed in marshes or even in small, tempo-rary water bodies, much vegetated. It was found onlyin two samples (2%) (Figs 3, 5).

AMIX – mixed assemblage. This fauna is composedof relatively equal proportions of the main ecologicalgroups. It was found in six samples (9%) (Figs 3, 5).

AO – assemblage with open-country species. It is afauna comprising two main components: snails ofrelatively dry, open habitats: Pupilla muscorum (Lin-naeus), Vallonia pulchella (O. F. Müller) and V. costata(O. F. Müller) and water molluscs represented mainlyby shells of Bithynella austriaca (Frauenfeld.). The re-maining ecological groups are represented by single


Fig. 5. Frequency of occurrence of mollusc assemblages. Forsymbols of assemblages see text

Fig. 4. Shade-lowing and open-country mollusc assemblagesof Upper Holocene calcareous tufa in Podhale Basin andPieniny Mts. Ecological groups of molluscs (according toLOZEK 1964, S. W. ALEXANDROWICZ 1987, 1999): M –mesophile species, O – open-country species, F – forestspecies. For symbols of assemblages see text


specimens or absent. This assemblage was found inone sample of calcareous tufa precipitated close to aspring within a completely deforested area (Figs 4, 5).

The diversity index TDI indicates mollusc diversityin particular samples. Its values vary between 0.07(poor communities dominated by one taxon) and0.99 (communities with nearly equal number of speci-mens of different species) (S. W. ALEXANDROWICZ1987, 1999). The relations between TDI and the num-ber of species allow to distinguish four types of assem-blages (Figs 6, 7). Poor assemblages (less than 10 spe-cies per sample) with low TDI values (<0.5) belong tothe first type (A). Oligomictic, less diversified assem-blages recognised in nine samples (12% of the totalnumber) can be assigned to this type (Figs 6, 7).Oligomictic, multispecies assemblages (B) character-ised by low TDI values (<0.5), but composed of morethan 10 taxa per sample, were found only in twosamples (3%) (Figs 6, 7). The third type comprises alimited number of taxa (less than 10) but is character-ised by relatively high TDI values, exceeding 0.5 (C).Such fauna was found in 27% of the analysed samples(20 samples) (Figs 6, 7). Rich and diverse assemblagesoccur in the majority of samples (42–58%) as themost common type of assemblages – D (Figs 6, 7).

The composition and structure of the assemblagesin the whole set of samples are described using indi-ces of constancy (C) and dominance (D). Only onespecies – Bithynella austriaca (Frauenfeld), can beplaced in the highest classes of C and D (C=5, D=5).This snail occurs most commonly (65 samples – 89%of the total) and is often represented by numerous

shells (3,279 specimens – 40% of the total). Six othertaxa: Aegopinella pura (Alder), Perforatella vicina(Rossmässler), Euconulus fulvus (O. F. Müller),Cochlicopa lubrica (O. F. Müller), Carychium minimumO. F. Müller and Galba truncatula (O. F. Müller) are in-cluded in classes C=3 and D=1, while the remainingspecies (67) are represented only by a low number ofspecimens, with C=2 or C=1, D=1 (Fig. 8).

The presented molluscan assemblages include spe-cies of various geographical range. The zoogeographi-cal structure of the fauna found in Late Holocene andrecently accumulated calcareous tufa comprises three


Fig. 6. Diversity of mollusc assemblages of Upper Holocenecalcareous tufa in Podhale Basin and Pieniny Mts. Typesof assemblages: 1 – oligomictic, low-diversity assemblages(A), 2 – oligomictic multiple assemblages (B), 3 – poli-mictic, low-diversity assemblages (C), 4 – polimicticmultiple assemblages (D); a log TDI – antilogarithmTDI; Ntax – number of taxa

Fig. 7. Proportion of types of mollusc assemblages in Pod-hale Basin and Pieniny Mts. A – oligomictic, low-diversityassemblages, B – oligomictic multiple assemblages, C –polymictic, low-diversity assemblages, D – polymicticmultiple assemblages

D-1 D-2 D-3 D-4 D-5

C-5

C-4

C-3

C-2

C-1

6

19

48

1

D=5; C=5 Bythinella austriaca

Aegopinella pura

Perforatella vicina

Cochlicopa lubrica

Euconulus fulvus

Carychium minimum

Galba truncatula

D=

1;

C=

3

D =1iC =7.75

i

Fig. 8. Constancy and dominance of assemblages of UpperHolocene calcareous tufa in Podhale Basin and PieninyMts. C – constancy, D – dominance, Ci – normalised in-dex of constancy (according to S. W. ALEXANDROWICZ1999), Di –– normalised index of dominance (accordingto S. W. ALEXANDROWICZ 1999)


main groups of taxa: widespeared species (HP), Cen-tral-European species (ME) and species of limitedgeographical range (EO) (Fig. 9). The first categoryincludes Holarctic (Hl), Palaearctic (Pl), European(Ep) and Eurosiberian (Es) taxa and constitutes 42%of the whole collected material (Fig. 9). Holarcticsnails are represented mainly by Cochlicopa lubrica (O.F. Müller), Euconulus fulvus (O. F. Müller) and Galba

truncatula (O. F. Müller) (14% of the fauna). Euro-pean taxa: Aegopinella pura (Alder) and Vitreacrystallina (O. F. Müller) constitute 12%, Eurosiberianspecies – Pisidium personatum Malm., Succinea oblongaDraparnaud and Carychium minimum O. F. Müller –11% of the fauna, while Palaearctic taxa: Punctumpygmaeum (Draparnaud) and Nesovitrea hammonis(Ström) are less numerous (5%).

Species inhabiting mainly Central Europe (ME)are the most important components of the assem-blages (54% of the analysed material). Among them,montane molluscs associated with the Carpathiansand the Alps (Ma): Bithynella austriaca (Frauenfeld),Vestia gulo (E. A. Bielz.) and Perforatella vicina(Rossmässler) occur very frequently, forming about49% of the fauna. The proportion of Central Euro-pean taxa (Me), such as Perforatella incarnata (O. F.Müller) and Arianta arbustorum (Linnaeus) is onlyabout 5%. Species of limited geographical range canbe regarded as subordinate components. They in-clude mainly South European (Em) species: Vitreadiaphana (Studer) and V. subrimata (Reinhardt) andform about 3% of the material. The remaining zoo-geographical groups: East European (Ee), Balkanand Pontic (Eb), West European (Ew) and Boreal(Ba) are represented only by single shells and in totalconstitute 1% of the described fauna (Fig. 9).

CONCLUSIONS

Late Holocene and recently precipitated calcareoustufa occur commonly in the Podhale Basin and in thePieniny Mts. In most outcrops and profiles presentedin this paper hard, porous travertines and granularcalcareous tufas are accessible. These latter depositsform layers and lenses, or fill niches inside the traver-tine covers. In several localities, fans built ofcoarse-grained calcareous tufa with numerous blocksof travertines and angular fragments of base rock arefound. Granular calcareous tufa usually containsmore or less rich and diverse molluscan assemblages.The described sediments are precipitated most oftenclose to springs, especially mineralised ones. Accord-ing to their lithological variability, three main typescan be distinguished. The first type includes typicalspring travertines in the form of covers or encru-stations on plants. The second type comprises granu-lar calcareous tufa with numerous shells of Bithynellaaustriaca (Frauenfeld), found only in small niches orfissures, as well as layers, lenses and fans built ofcoarse-grained calcareous tufa, usually containingblocks of travertines, pebbles and angular fragmentsof base rock, deposited on terraces. They often con-tain rich and diverse mollusc assemblages with a highproportion of shade-loving species, catholic speciesand numerous shells of B. austriaca. The last type is

represented by fine-grained calcareous tufa, depos-ited in temporary, small and much vegetated waterbodies developed close to springs. Water species,mainly B. austriaca and Galba truncatula (O. F.Müller), accompanied by a hygrophile snail – Succineaputris (Linnaeus) and taxa inhabiting moist forests,like Perforatella vicina (Rossmässler), are the most im-portant components of the fauna.

Recently precipitated calcareous tufas and traverti-nes, mainly travertine covers, are often partly or evencompletely destroyed during floods. However, they re-generate in the same places during several years (MA-STELLA 1975, GRUSZCZYÑSKI & MASTELLA 1986) andrepresent only a very short time span. Calcareoustufas deposited on terraces in small water bodies aremore stable and correspond with much longer peri-ods (several dozen or even several hundred years).

Malacological analyses made it possible to distin-guished seven types and two sub-types of mollusc as-semblages. Communities with a high proportion ofshade-loving species occur commonly. However, thefauna with Bithynella austriaca (Frauenfeld) seems tobe the most characteristic of recently precipitated cal-careous tufas. Particular assemblages are strongly as-sociated with local environmental conditions.Polymictic communities are the most common in the


Fig. 9. Zoogeographical structure of molluscan fauna fromUpper Holocene calcareous tufa in Podhale Basin andPieniny Mts. HP – widespread species, ME – Middle-Eu-ropean species, EO – European species of limited ran


analysed material. Similar, strongly differentiated fau-nas have been described from numerous localities ofLate Glacial and Holocene calcareous tufa in the Pol-ish Carpathians (S. W. ALEXANDROWICZ 1984b, 1994,S. W. ALEXANDROWICZ & Z. ALEXANDROWICZ 1998,1999, S. W. ALEXANDROWICZ & GERLACH 1983, S. W.ALEXANDROWICZ & CHMIELOWIEC 1991, W. P.ALEXANDROWICZ 1997, 2001, 2003, 2004) and in Mid-dle Polish Uplands (S. W. ALEXANDROWICZ 1983, W.P. ALEXANDROWICZ 2004).

The poor assemblages in several samples are domi-nated by Bithynella austriaca (Frauenfeld). Such fau-nas have been recorded only from the Carpathians (S.W. ALEXANDROWICZ 1984b, W. P. ALEXANDROWICZ1997, 2004). B. austriaca is the most important speciesfor recently precipitated calcareous tufa and was iden-tified in the majority of samples, often in large num-bers. In calcareous tufas of Late Glacial and Holoceneage the species is very rare or absent (S. W.ALEXANDROWICZ 1983, W. P. ALEXANDROWICZ 2004).This problem is difficult to explain directly. Shells ofB. austriaca are thin and very delicate, and prone tophysical and chemical destruction. On the other

hand, they occur in large numbers only in profiles oftypical spring travertines, where granular calcareoustufa occurs in small niches or fissures and can be eas-ily removed by erosion.

Malacological investigations of recently precipi-tated calcareous tufas are very important for the inter-pretation of subfossil assemblages found in depositsdeveloped of the Late Glacial and Holocene. Calcare-ous sediments accumulated on the surface of low ter-races, forming fans on relatively flat slopes or fillingsmall water bodies can be preserved for long periods(hundreds or even thousands of years). Deposits ac-cumulated close to springs are usually not older thanseveral years. From this point of view, the mass occur-rence of Bithynella austriaca (Frauenfeld), typical ofthe last-mentioned sediments, can be regarded as anindicator of their age.

ACKNOWLEDGEMENTS

This study was sponsored by AGH University of Sci-ence and Technology through the University grant no11.11.140.560.

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Received: December 15th, 2009Accepted: June 1st, 2010



MOLLUSCAN ASSEMBLAGES OF RECENT CALCAREOUS TUFAS IN THE PODHALE BASIN AND PIENINY MTS (S. POLAND)

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